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Cactaceae, Cactoideae) Botanical Sciences 90 (2): 97-108, 2012 BOTÁNICA ESTRUCTURAL EL HÁBITO Y LA FORMA DE CRECIMIENTO EN LA TRIBU CACTEAE (CACTACEAE, CACTOIDEAE) MONSERRAT VÁZQUEZ-SÁNCHEZ1, 3, TERESA TERRAZAS1 Y SALVADOR ARIAS2 1Departamento de Botánica, Instituto de Biología, Universidad Nacional Autónoma de México 2Jardín Botánico, Instituto de Biología, Universidad Nacional Autónoma de México 3Autor para la correspondencia: [email protected] Resumen: Los términos ‘forma de crecimiento’ y ‘hábito’ con frecuencia se usan como sinónimos; sin embargo, su asignación a las diferentes especies resulta problemática en algunos grupos taxonómicos debido a la diversidad morfológica; éste es el caso en Cactaceae, particularmente en la tribu Cacteae de la subfamilia Cactoideae. Se estudió la morfología del tallo de 102 especies (26 géneros) de Cacteae con el objetivo de identificar el hábito y reconocer cuántas formas de crecimiento se presentan en la tribu, así como para hacer una distinción clara entre los términos ‘hábito’ y ‘forma de crecimiento’ para Cactaceae y discutir sus diferencias respecto a los conceptos de forma de vida y arquitectura vegetal. Con base en las observaciones y mediciones para 102 especies de Cacteae se reconocieron cuatro formas de crecimiento (cilíndrica, columnar, globosa y globosa-deprimida). Por su tamaño y lon- gevidad no es posible utilizar los términos ‘árbol’ o ‘hierba’ para las especies de Cacteae. El 12% de las especies estudiadas de esta tribu tienen ramificación basítona (arbustos) comoAcharagma roseana, Ferocactus pilosus y Thelocactus bicolor. Sugerimos utili- zar únicamente el término ‘forma crecimiento’ para referirse a los tallos de la tribu Cacteae con la finalidad de evitar confusiones. Palabras clave: arbusto, arquitectura vegetal, cilíndrica, columnar, forma de vida, globoso. Abstract: The terms ‘growth form’ and ‘habit’ are often used as synonyms. However, their assignment to different species is problematic because of the morphological diversity occurring in some taxonomic groups, as is the case in Cactaceae, particularly in the tribe Cacteae of the Cactoideae subfamily. Stem morphology was studied in 102 species (26 genera) of Cacteae in order to identify the habit and to recognize how many growth forms occur in the tribe, as well to make a distinction between the concepts of habit and growth form in Cactaceae, and to discuss the differences between them and the concepts of life form and plant archi- tecture. Based on observations and measurements for 102 species of Cacteae, four growth forms (cylindrical, columnar, globose, and globose-depressed) were recognized. Neither the habit ‘tree’ or ‘herb’ can be assigned to any member of Cacteae based on their size or longevity. Twelve percent of the studied species of this tribe have basitonic branching (shrubs) as for Acharagma roseana, Ferocactus pilosus and Thelocactus bicolor. In order to avoid confusion, we suggest using only the term ‘growth form’ when refe- rring to the various stem forms in the Cacteae tribe. Keywords: columnar, cylindric, globose, life form, plant architecture, shrub. l término hábito, del latín habitus, significa porte o as- ‘Forma de crecimiento’ y ‘forma de vida’ se usan fre- Epecto exterior (Font Quer, 1977). Harris y Harris (2001) cuentemente como sinónimos de ‘hábito’ al referirse a la definen ‘hábito’ como la apariencia general o el modo de forma de una planta (Mueller-Dombois y Ellenberg, 1974; crecimiento de una planta. Para Judd et al. (2002) este tér- Moreno, 1984; Pavón et al., 2000). Sin embargo, el término mino denota la apariencia general de una planta y conside- ‘forma de vida’ fue propuesto por Raunkiaer (1934) para ran tres hábitos: (1) árbol (con un tronco bien definido), (2) clasificar a las plantas con base en la posición de las yemas arbusto (varias ramas que surgen en la posición basal y más u órganos de renuevo, a partir de los cuales se desarrollarán bajo que un árbol), ambos con tejido leñoso (lignificados), nuevos brotes o nuevo follaje después de la época desfa- y (3) hierba (tallos anuales no lignificados). Los términos vorable. Ejemplos de formas de vida según Raunkiaer son ‘árbol’, ‘arbusto’ y ‘hierba’ son el antecedente más antiguo ‘criptófita’ y ‘geófita’, y estos términos tienen una conno- de clasificación propuesto por Teofrasto (300 A.C.) con base tación adaptativo-ecológica (Mueller-Dombois y Ellenberg, en la forma de las plantas (Díaz et al., 2002). 1974). Por el contrario, ‘forma de crecimiento’ describe el 97 MONSERRAT VÁZQUEZ-SÁNCHEZ ET AL. diseño o figura y la construcción de los individuos, junto mientras que cuando la ramificación es basítona correspon- con los cambios que sufre a lo largo de su vida. Algunos au- derían a arbustos. tores, además de incluir en ella a las hierbas, los arbustos y En plantas de hábito arbustivo todas las ramas surgen los árboles, mencionan a las enredaderas y a las lianas, o bien cerca del nivel del suelo (ramificación basítona) como en únicamente se refieren a plantas erectas, rastreras o trepado- Leptocereus quadricostatus (Bello) Britton & Rose, Neorai- ras. De acuerdo con el uso actual del lenguaje, los conceptos mondia arequipensis (Meyen) Backeb. o Stenocereus stella- de hábito y de forma de crecimiento no pueden separarse en tus (Pfeiff.) Riccob. (Figura 1G). En las plantas con hábito la mayoría de los grupos taxonómicos. Un concepto más am- arbustivo las ramas pueden ser decumbentes, como en Pe- plio es el de ‘arquitectura vegetal’, el cual considera cuatro niocereus greggii (Engelm.) Britton & Rose o S. gummosus aspectos con base en el reconocimiento de las unidades de (Engelm. ex Brandegee) A.C.Gibson & K.E.Horak (Figura construcción: el tipo de crecimiento (rítmico o continuo), el 1H); postradas, como en Haageocereus albispinus (Akers) patrón de ramificación (monopodial o simpodial), la diferen- Backeb. y S. eruca (Brandegee) A.C.Gibson & K.E.Horak ciación morfológica de los ejes (ortótropa o plagiótropa) y la (Figura 1I), especie cuyas ramas parecen desplazarse por posición de las estructuras sexuales (terminal o lateral) para el suelo (Bravo-Hollis, 1978; Anderson, 2001); o péndu- asignar un modelo (Barthélémy y Caraglio, 2007; Bell y Br- las, como en algunas especies de Disocactus y Selenicereus yan, 2008). Estos atributos del estudio arquitectónico pueden (Anderson, 2001). Cuando en la familia Cactaceae el hábi- ser aplicados a cualquier planta, ya sea que se trate de un ár- to es arborescente o arbustivo, como se ejemplificó en los bol, un arbusto, una hierba o una liana (Hallé et al., 1978). párrafos anteriores, la asignación de estos términos no es Los miembros de la familia Cactaceae se caracterizan por difícil; sin embargo, hay tribus como Cacteae donde la asig- la morfología peculiar de sus tallos. El hábito en la familia nación del hábito no es fácil. es referido como hábito de crecimiento (Kaplan, 2001) o En la tribu Cacteae, sujeto de este estudio, predominan forma de crecimiento (Buxbaum, 1950; Bravo-Hollis, 1978; los géneros con individuos de talla baja, como Aztekium y Rauh, 1979; Anderson, 2001). En sentido estricto se pue- Mammillaria, pero también hay géneros cuyos integrantes de considerar que en la familia Cactaceae hay dos hábitos: tienen alturas de más de 1 m, como Echinocactus y Fero- arbóreo y arbustivo. Por ejemplo, en la subfamilia Cactoi- cactus, o son de tallas intermedias (20-30 cm), como Es- deae se presentan las modificaciones más evidentes de los cobaria, Neolloydia y Thelocactus. ¿Podrían aplicarse los tallos; cuando el hábito corresponde a un árbol sin ramas es conceptos de hábito arbóreo (árbol) y herbáceo (hierba) a los llamado columnar y cuando se ramifica se le conoce como miembros de la tribu?, ¿son arbustos algunos miembros de candelabriforme. Cuando la planta se ramifica se distingue los géneros Acharagma, Mammillaria y Thelocactus? An- en ella un tronco con ramas ortotrópicas (verticales) y pa- derson (2001) señala que en la tribu Cacteae las formas de ralelas entre sí, como en Pachycereus weberi (J.M.Coult.) crecimiento son cespitosas, globosas, globoso-deprimidas o Backeb. (Figura 1A) o Stenocereus dumortieri (Scheidw.) con forma de barril. Algunas de las formas descritas por An- Backeb.; la ramificación se simplifica, como en Carnegiea derson (2001) no podrían ser aplicadas en sentido estricto gigantea (Engelm.) Britton & Rose (Figura 1B), hasta des- a la forma de crecimiento en una cactácea; por ejemplo, la aparecer, como en Cephalocereus columna-trajani (Karw. forma de crecimiento cespitosa. El término ‘cespitoso’ tam- ex Pfeiff.) K.Schum. y Stephanocereus leucostele (Gürke) bién se ha utilizado para otras familias como Aizoaceae y A.Berg. (Figura 1C). En estas dos últimas especies se ad- Crassulaceae (Eggli y Newton, 2004) con tallos suculentos. quiere la forma de crecimiento columnar (Buxbaum, 1950; Este término entonces se refiere a la ramificación pero no Bravo-Hollis, 1978; Anderson, 2001), pero el hábito sigue a la forma de crecimiento, como puede notarse en las des- siendo arbóreo. cripciones de Benson (1982), quien no reconoce la forma de El tipo de ramificación en Cactaceae puede ser acróto- crecimiento cespitosa. El otro término empleado en la litera- na cuando las ramas aparecen cerca del ápice del tronco tura de cactáceas se refiere a la forma de crecimiento que en principal, como sucede en Browningia candelaris (Meyen) términos coloquiales se define como barril (Bravo-Hollis, Britton & Rose (Figura 1D; Bravo-Hollis, 1978); dicóto- 1978; Gibson y Nobel, 1986; Bravo-Hollis y Sánchez-Me- ma cuando el meristemo apical se divide en dos, como en jorada, 1991; Anderson, 2001), el cual alude a una forma Hatiora y Rhipsalis (Anderson, 2001); mesótona si de un cilíndrica con las variantes subcilíndrica o ligeramente ci- tronco bien definido salen ramas que se van ramificando su- líndrica. ¿Cuáles serían los términos a utilizar para evitar cesivamente hasta formar una copa amplia, como en Myr- ambigüedad? Entonces, las preguntas a responder en este tillocactus geometrizans (Mart. ex Pfeiff.) Console (Figura estudio fueron: ¿cuántas formas de crecimiento hay en Cac- 1E), Polaskia chichipe (Gosselin) Backeb.
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